Kenophytes in Different Forest Types of Ukraine

18/2 • 2019, 289–312 DOI: 10.2478/hacq-2018-0013

Kenophytes in different forest types of Ukraine

Viktor Onyshchenko1 

Key words: forest, vegetation, Abstract alien plants, kenophytes, invasion, The article aims to 1) describe the distribution of non-native species of vascular habitats. plants (only kenophytes, i. e. naturalized species introduced after the 15th century) in different types of forests and different biogeographical regions of Ukraine; Ključne besede: gozd, vegetacija, 2) compare proportions of kenophytes in forests of different areas; 3) detect statis- tujerodne rastline, kenofiti, invazija, tically significant changes in the occurrence of kenophytes over the last 80 years. habitati. The material consists of 2701 relevés sampled in 1990‒2018. They were taken from Ukrainian phytocoenological publications and databases. In Ukraine, as in other European countries, the highest proportion of kenophytes (percentage of species number per relevé) is in floodplain forests (up to 9.1% in willow and pop- lar forests). The lowest proportion is characteristic for bog woods (0%) and most types of mountain forests. Among biogeographical regions, the highest values were found in the flatland areas of the Steppic region and the forest-steppe part of the Continental region. The most frequent species are Impatiens parviflora (predominantly in broadleaved woods, absent in relevés from the steppic biogeoregion) and Erigeron canadensis (pine woods on sand). Comparison with 1466 older relevés sampled in 1950–1989 allows us to make a conclusion that the proportion of kenophytes increased at least in one habitat type (oak-hornbeam forests).

Izvleček V članku smo želeli 1) opisati razširjenost tujerodnih rastlinskih vrst (samo kenofitov, to je naturaliziranih vrst, ki so se pojavile po 15. stoletju) v različnih gozdnih habitatnih tipih in v različnih biogeografskih regijah v Ukrajini; 2) primerjati deleže kenofitov v gozdovih v različnih območjih; 3) prikazati statistično značilne spremembe v pojavljanju kenofitov v zadnjih 80. letih. Podatke predstavlja 2701 popis, ki so bili narejeni v obdobju med 1990 in 2018. Uporabili smo fitoceno- loško literaturo in podatkovne baze iz Ukrajine. V Ukrajini, kot tudi v ostalih državah v Evropi, je najvišji delež kenofitov (odstotek vrst na popis) v poplavnih gozdovih (do 9,1% v vrbovih in topolovih gozdovih). Najnižji odstotek je značilen za barjanske gozdove (0%) in večino habitatnih tipov gorskih gozdov. Med bioge- ografskimi regijami je najvišji delež v nižinskih predelih stepske regije in gozdne stepe v kontinentalni regiji. Najbolj pogosti vrsti sta Impatiens parviflora(prevla - duje v listopadnih gozdovih, odsotna je v popisih iz stepske biogeografske regije) Received: 13. 4. 2018 in Erigeron canadensis (v borovih gozdovih na pesku). S primerjavo 1466 starejših Revision received: 4. 12. 2018 popisov, narejenih med leti 1950 in 1989 smo prišli do zaključka, da se je delež Accepted: 5. 1. 2019 kenofitov povečal vsaj v enem habitatnem tipu (hrastovo-gabrovi gozdovi).

1 M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Ukraine. E‑mail: [email protected]

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ropean Russia occupy 49% of the study area but they are Introduction represented only by 1% of the total number of relevés. Phytosociological relevés are an important source of in- We made the assessment of occurrences of kenophytes in formation about ecological preferences of species, their forests of Ukraine using a higher number of relevés. A geographical distribution and dynamics. In recent decades more narrow geographical scope allows using the results some studies were published that show geographical and for comparisons between regions of Europe. The analy- ecological patterns of alien species on the basis of phyto- sis for the territory of Ukraine, even without compari- coenological data. The study areas were the Czech Repub- son with other countries, may reveal some geographical lic (Chytrý at al. 2005), Catalonia (Vila et al. 2007), the patterns because Ukraine comprises parts of four biogeo- Basque Country (Campos et al. 2013), Slovakia (Med- graphical regions and measures about 1300 km in length. vecká et al. 2014) and Slovenia (Küzmič & Šilc 2017). At present there is no information in literature about the Chytrý et al. (2008) performed a comparison of plant level of invasion in forests of the Steppic biogeographical invasions for three areas of Europe (the Czech Republic, region of Europe. In other bioregions of Europe, estima- Catalonia, Great Britain) that represent three biogeo- tions were carried out only at higher levels of habitat clas- graphical regions (Continental, Mediterranean, Atlantic). sifications. Estimations in the aforementioned articles were made for Presence of 40 alien invasive species within plant com- classes and alliances of vegetation or for EUNIS level 2 munities of Ukraine at the level of alliances of vegetation habitats (Davies et al 2004, EUNIS habitat …). Accord- without quantitative estimations was described in article ing to these studies the lowest proportions of aliens in by Abduloyeva & Karpenko (2009). all regions are in bogs, mires, alpine and subalpine grass- The aim of the present study is to describe the frequency lands, heaths on poor soils. The highest proportions are of kenophytes in forest types and different biogeographi- found in synanthropic vegetation, marine coastal habitats, cal regions of Ukraine, and to compare the results with standing inland waters (only kenophytes), sedge and reed published data on the subject from other areas. beds, riverine shrubs. Wagner et al. (2017) published the assessment of plant invasions in European forests that is based on the analy- Materials and methods sis of 83,396 relevés from the European Vegetation Ar- The study area includes the entire territory of Ukraine. chive (Chytrý at al. 2016) sampled in 1970–2015. The According to the data on the European Environment assessment was made for habitat types of the revised Agency website (Europe’s biodiversity ... 2018) Ukraine version of EUNIS habitat classification (Schaminée et al. comprises parts of four European biogeographical re- 2014). These habitat types approximately correspond to gions: Alpine, Continental, Pannonian and Steppic. Two level 3 or 4 habitats of the official EUNIS habitat classifi- regions (Continental and Steppic) were subdivided ac- cation. All analyses were performed only for kenophytes cording to the geobotanical subdivision of Ukraine in the (naturalized species introduced voluntarily or involun- National Atlas of Ukraine (2008) (Figure 1). tarily after the 15th century). Archaeophytes (introduced The relevés for the analysis were taken from the data- before the end of the 15th century) were excluded from base EU-UA-006 “Vegetation Database of Ukraine and consideration because the status of archaeophytes is poorly known in some countries. The highest proportion of kenophytes was found in riparian woods. The lowest proportions were in mountain forests and mire woodlands. The most frequent kenophytes of European woods turned out to be Impatiens parviflora (21.1% of total frequency of kenophytes), Padus serotina (8.8%), Robinia pseudoacacia (7.5%). The highest number of occurrences of kenophytes belongs to phanerophytes and therophytes (for native species hemicryptophytes and phanerophytes). Species originating in Europe constitute 47% of the total number of kenophytes in European woods, and their total frequency is 22% of total frequency of kenophytes. The distribution of relevés used in the article by Wag- ner et al. (2017) has a strong geographic bias: Ukraine, Figure 1: Biogeographical regions of Ukraine. Belarus, Moldova, Georgia, Armenia, Azerbaijan and Eu- Slika 1: Biogeografske regije Ukrajine.

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Adjacent Parts of Russia”, unpublished database of rele- of their natural range or in clearly unnatural conditions vés from the National Nature Park “Holosiivskyi”(Kyiv and natural woods of non-native species), 2) relevés that city) and numerous Ukrainian publications (Andrienko have no species data obtained in period May 15 – Au- et al. 2004, Bairak 1996, Budzhak & Onyshchenko gust 31, 3) relevés that could not be unequivocally as- 2004, Chorney et al. 2005, Derzhypilsky et al. 2011, signed to either habitat type. The area of the relevés varies Didukh 1996, 2003, Didukh et al. 2003, Fitsailo 2003, from 100 to 2500 m2, predominantly 400–900 m2. This Gomlya 2005, Goncharenko 2001, 2003, Klimuk et al. parameter was not used in relevé filtering. Some analy- 2006, Konishchuk 2003, Kuzemko 2001, Kuzemko & ses showed that the effect of plot size on the proportion Chorna 2002, Lukash & Onyshchenko 2006, Melnyk & of species is negligible (Chytrý et al. 2005, Vilà et al. Korinko 2005, Onyshchenko 1998, 2002, 2007, 2009, 2007). The final number of relevés was 2701. In order to 2013a, 2013b, Onyshchenko & Budzhak 2003, Onysh- compare the current frequency of kenophytes with their chenko &t Lubinska 2006, Onyshchenko & Lukash frequency in previous decades 1466 relevés sampled in 2004, Onyshchenko & Sidenko 2002, Onyshchenko & 1950–1989 were used (Bukhalo 1962, Grygora et al. Yuglichek 2010, Onyshchenko et al. 2007, 2009, 2015, 2005, Vorobyov et al. 1997, Vorobyov et al. 2008). Old 2016, Orlov & Yakushenko 2005, Panchenko 2013, relevés were not sampled in the same areas as the relevés Panchenko & Onyshchenko 2003, Ralo & Onyshchen- from 1990–2018. This reduces the possibility of com- ko 2008, Shelyag-Sosonko et al. 2003, Shevchyk et al. parison of two sets of data. 1996a, 1996b, 1997, I. Solomakha et al. 2006, 2015, The names of species follow the database Euro+Med V. Solomakha et al. 2004, 2016, Soroka 2008, Vorobyov (2006–2018). Residence statuses of species (whether a et al. 2002, Yakushenko 2003, Yuglichek & Onyshchen- species is a kenophyte or not) were taken from Ukrainian ko 2003). All relevés selected for estimation of modern publications (Protopopova 1991, Mosyakin & Yavorska occurrence of kenophytes were sampled in 1990–2018. 2003, Bagrikova 2013, Solomakha et al. 2015). Some Each relevé was assigned to a habitat of the revised ver- species are kenophytes only in a part of their distribution sion of the EUNIS habitat classification (European Red area in Ukraine (Acer pseudoplatanus, Arrhenatherum List of Habitats, Schaminée et al. 2014) (Table 1). Some elatuis, Berberis vulgaris, Corynephorus canescens, Fagus relevés were excluded from the analysis: 1) relevés repre- sylvatica, Ligustrum vulgare, Picea abies, Spergula moriso- senting highly artificial woods (plantations of species out nii, Vinca minor). They were involved into the analysis

Table 1: Codes, names of habitats and number of relevés. Tabela 1: Šifra, ime habitanega tipa in število popisov.

Habitat Number of relevés Number of Habitat name code before 1990 relevés since 1990 G1.1 Temperate and boreal softwood riparian woodland - 84 G1.2a Alnus woodland on riparian and upland soils - 93 G1.2b Temperate and boreal hardwood riparian woodland - 95 G1.4 Broadleaved swamp woodland on non-acid peat 215 85 G1.5 Broadleaved bog woodland on acid peat 316 - G1.6a Fagus woodland on non-acid soils - 459 G1.6b Fagus woodland on acid soils - 37 G1.7a Temperate and submediterranean thermophilous deciduous woodland 66 151 G1.8 Acidophilous Quercus woodland - 26 G1.Aa Carpinus and Quercus mesic deciduous woodland 630 911 G1.Ab Ravine woodland - 90 G3.1a Temperate mountain Picea woodland - 121 G3.1b Temperate mountain Abies woodland - 44 G3.2 Temperate subalpine Larix, Pinus cembra and Pinus uncinata woodland - 5 G3.4a Temperate and continental Pinus sylvestris woodland 120 388 G3.4b Temperate and submediterranean montane Pinus sylvestris-Pinus nigra woodland 80 17 G3.A Picea taiga woodland 27 17 G3.Da Pinus mire woodland 44 78 All habitats 1498 2701

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only in the cases when they met outside their old dis- Quercus rubra (7.9%), Acer negundo (7.7%), Erigeron tribution area. For each of the species its constancy in annuus (3.8%), Robinia pseudoacacia (3.4%). Impatiens every habitat type, constancy in total set of relevés, con- parviflora is the most frequent kenophyte in the Conti- stancy in Ukrainian forests (mean for habitats types with nental, the Alpine and the Pannonian biogeographical weighting proportionally to habitat area) and proportion regions. It is not present in forest relevés from the Steppic of species occurrences in the sum of occurrences of all biogeographical region. Erigeron annuus is present only kenophytes were calculated. in relevés from the Continental and the Alpine biogeographical regions. The largest average proportion of kenophytes in relevés Results (9.1%) is peculiar to G1.1 “Floodplain Salix- and Populus- Selected relevés were referred to 17 habitat types of the dominated woods” (Figure 2). Other habitats with high modified EUNIS classification (Table 1). Mean propor- proportions of kenophytes are G1.2a “Alnus woodland on tion of kenophytes for all relevés is 1.13%, mean weight- riparian and upland soils” (2.3%) and G1.2b “Temperate ed proportionally to habitat areas is 1.02%. A total of and boreal hardwood riparian woodland” (3.1%). Keno- 62 kenophytes are present in the dataset. Their distribu- phytes are not present in relevés of coniferous bog wood- tion along habitat types is shown in Table 2. The most lands (G3.Da), acidophilous beech forests (G1.6b), frequent kenophytes in Ukrainian woods are Impatiens mountain pine forests (G3.2, G3.4b). A very low propor- parviflora (20.6% of the sum of kenophytes constancies), tion is in G3.1a “Temperate mountain Picea woodland” Erigeron canadensis (14.8%), Prunus serotina (9.4%), (0.03%). Other habitats have intermediate values of this

Table 2: Constancy of kenophyte species in habitat types of Ukraine (habitats types without kenophytes were excluded). Tabela 2: Stalnost kenofitov v habitatnih tipih v Ukrajini (habitatni tipi brez prisotnih kenofitov niso prikazani). G1.1 G1.2a G1.2b G1.4 G1.6a G1.7a G1.Aa G1.Ab G3.1a G3.1b G3.4a Constancy in dataset (%) ‘% of total kenophytes’ occurrences Constancy with of plots weighting proportionally to (%) habitat areas Number of relevés 84 93 95 85 459 151 911 90 121 44 388 Number of kenophytes species

Acer negundo 39.3 2.2 16.8 11.8 . 4.0 2.4 . . . 1.0 3.41 11.7 2.35 Acer pseudoplatanus* . 1.1 3.2 1.2 . . 0.4 . . . . 0.33 1.1 0.27 Acer saccharinum ...... 0.3 0.04 0.1 0.09 Aesculus hippocastanum . 1.1 . . . . 0.2 . . . . 0.11 0.4 0.09 Ambrosia artemisifolia ...... 2.1 0.30 1.0 0.73 Amelanchier ovalis . . 3.2 ...... 0.3 0.15 0.5 0.21 Amelanchier spicata . . 1.1 . . 6.0 . . . . 1.3 0.56 1.9 0.65 Amorpha fruticosa 35.7 . . 1.2 ...... 0.5 1.22 4.2 0.65 Armeniaca vulgaris 1.2 6.5 ...... 0.26 0.9 0.27 Arrhenatherum elatius* . . . . . 2.0 . . . . . 0.11 0.4 0.05 Berberis vulgaris* . . . . . 2.0 . . . . 1.5 0.33 1.1 0.60 Bidens frondosus 16.7 6.5 1.1 7.1 ...... 1.00 3.4 0.55 Bryonia alba 1.2 ...... 0.04 0.1 0.02 Cannabis sativa . . 1.1 . . 0.7 . . . . . 0.07 0.3 0.06 Caragana arborescens . 1.1 2.1 . . 1.3 0.7 . . . 0.3 0.45 1.5 0.40 Corynephorus canescens* ...... 1.5 0.22 0.8 0.54 Cuscuta campestris ...... 0.3 0.04 0.1 0.09 Digitaria aegyptiaca ...... 0.3 0.04 0.1 0.09 Echinocystis lobata 3.6 . . 4.7 ...... 0.26 0.9 0.07 Eragrostis minor ...... 1.3 0.19 0.6 0.45

292 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine G1.1 G1.2a G1.2b G1.4 G1.6a G1.7a G1.Aa G1.Ab G3.1a G3.1b G3.4a Constancy in dataset (%) ‘% of total kenophytes’ occurrences Constancy with of plots weighting proportionally to (%) habitat areas Number of relevés 84 93 95 85 459 151 911 90 121 44 388 Erigeron annuus 7.1 . 2.1 2.4 0.4 2.6 1.6 2.2 . . 0.8 1.33 4.6 0.96 Erigeron canadensis 13.1 4.3 2.1 . . 5.3 . 1.2 . . 9.0 2.26 7.7 3.74 Fagus sylvatica* ...... 0.2 . . . . 0.07 0.3 0.05 Fraxinus americana 1.2 ...... 0.04 0.1 0.02 Gleditsia triacanthos . . . . . 0.7 0.2 . . . . 0.11 0.4 0.07 Hemerocallis lilioasphodelus 1.2 ...... 0.04 0.1 0.02 Hordeum murinum s. l. . 1.1 ...... 0.04 0.1 0.04 Hyoscyamus niger . . . . 1.1 6.0 . . . . . 0.52 1.8 0.25 Impatiens glandulifera . 1.1 ...... 0.04 0.1 0.04 Impatiens parviflora 3.6 21.5 46.3 15.3 5.9 18.5 8.2 16.7 0.8 4.5 0.8 8.55 29.3 6.11 Iva xanthiifolia . 1.1 ...... 0.04 0.1 0.04 Ligustrum vulgare* . . . . . 0.7 . . . . . 0.04 0.1 0.02 Lonicera tatarica . . . . . 2.6 0.1 . . . . 0.19 0.6 0.09 Morus alba 13.1 . . . . 0.7 0.2 . . . . 0.52 1.8 0.24 Morus nigra 2.4 ...... 0.07 0.3 0.03 Oenothera biennis 8.3 ...... 1.3 0.44 1.5 0.57 Oenothera rubricaulis . . . . . 0.7 . . . . 0.3 0.07 0.3 0.11 Oxalis dillenii 2.4 ...... 0.07 0.3 0.03 Oxalis stricta ...... 0.1 . . . . 0.04 0.1 0.03 Partenocissus quinquefolia 1.2 . . . . . 0.1 . . . . 0.07 0.3 0.04 Parthenocissus inserta . . 2.1 ...... 0.3 0.11 0.4 0.17 Physocarpus opulifolius ...... 0.5 0.07 0.3 0.18 Picea abies* ...... 0.7 . . . . 0.22 0.8 0.15 Prunus cerasifera . . . . . 2.0 . . . . . 0.11 0.4 0.05 Prunus serotina . 1.1 2.1 . . 4.6 0.2 . . . 5.4 1.22 4.2 2.20 Quercus rubra . 2.2 2.1 . . 1.3 2.3 1.2 . . 0.8 1.15 3.9 1.01 Robinia pseudoacacia 1.2 3.2 2.1 . 0.2 2.0 1.4 1.2 . 2.3 2.1 1.22 4.2 1.39 Saponaria officinalis . . . . . 2.6 . . . . . 0.15 0.5 0.07 Secale sylvestre 1.2 ...... 0.04 0.1 0.02 Senecio viscosus ...... 0.5 0.07 0.3 0.18 Sorbus intermedia . . . . . 0.7 . . . . . 0.04 0.1 0.02 Spergula morisonii* ...... 0.5 0.07 0.3 0.18 Spiraea chamaedryfolia ...... 1.2 . . . 0.04 0.1 0.00 Swida alba 1.2 . . 1.2 0.2 ...... 0.11 0.4 0.04 Symphoricarpos albus ...... 0.5 0.07 0.3 0.18 Triticum aestivum . . 1.1 ...... 0.04 0.1 0.04 Ulmus pumila 1.2 ...... 0.04 0.1 0.02 Veronica hederifolia . . . . . 2.6 0.3 . . . . 0.26 0.9 0.15 Vinca minor* . 1.1 1.1 . . . 0.4 . . . 0.3 0.26 0.9 0.28 Vitis vinifera 1.2 ...... 0.04 0.1 0.02 Xanthium orientale subsp. 1.2 ...... 0.1 0.02 riparium 0.04 Xanthium rupicola 2.4 ...... 0.07 0.3 0.03 Xanthium strumarium 2.4 ...... 0.07 0.3 0.03

* – species is native in a part of the Ukrainian territory.

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Figure 2: Mean proportions of kenophytes in relevés of different habitats in Ukraine (with standard deviation) and Europe. Data on Europe are from the article by Wagner et al. (2017). Slika 2: Povprečni deleži kenofitov v popisih v različnih habitatnih tipih v Ukrajini (s standardnim odklonom) in v Evropi. Podatki o pojavljanju v Evropi so iz članka Wagner et al. (2017).

Figure 3: Mean total covers of kenophytes in relevés of different habitats in Ukraine (relevés without kenophytes were included in the calculation with cover of kenophytes = 0). Slika 3: Povrečna skupna pokrovnost kenofitov v popisih v različnih habitatnih tipih v Ukrajini (popisi brez kenofitov so bili vključeni v izračun s pokrovnostjo kenofitov=0).

index. Proportions of kenophytes in habitats in different serotina. The most frequent kenophytes of habitat G3.4a areas are presented in Figures 4–6. “Temperate and continental Pinus sylvestris woodland” Major kenophytes of habitat G1.1 are Acer negundo are Erigeron canadensis (9%) and Prunus serotina (5%, (constancy 39%), Amorpha fruticosa (36%), Bidens fron- only in relevés from Kyiv). dosus (17%). These species often have high cover (Fig- The relevés from mountain regions have the smallest ure 3). Mean cover of Acer negundo in relevés of G1.1 proportion of kenophytes (Table 3). In the flatland part with the presence of the species is 8%, Acer negundo – of Ukraine, higher proportions are characteristic of the 7%, Bidens frondosus – 3%. These species are frequent Steppic biogeographical region and the forest-steppe part also in other types of hygrophilous broadleaved forests of the Continental biogeographical region. (G1.2a, G1.2b), however the most frequent kenophyte in these habitats is Impatiens parviflora (constancy 22% Table 3: Mean proportion of kenophytes in plots from different and 46%, mean cover 20% and 18% respectively). The biogeographical regions of Ukraine. typical habitats of G1.4 “Broadleaved swamp woodland Tabela 3: Povprečni delež kenofitov na popisnih ploskvah v on non-acid peat” are probably poor in kenophytes. The različnih biogeografskih regijah Ukrajine. available relevés represent mainly forests that are transi- Mean % of Number tional to G1.2a. The most frequent kenophyte of mesic Biogeographical region eutrophic broadleaved forests (G1.6a, G1.Aa, G1.Ab) is kenophytes in relevé of relevés Impatiens parviflora (constancy in these habitats 5%, 8%, Continental: deciduous forest 0.53 1276 17%, mean cover 0.1%, 7%, 1% respectively). In habitat Continental: forest-steppe 2.46 657 G1.7a “Temperate and submediterranean thermophilous Pannonian 1.02 26 deciduous woodland”, Impatiens parviflora also holds the Alpine 0.31 530 first position (constancy 19%, mean cover 4%). In addi- Steppic: flatland 3.13 180 tion, this habitat has significant constancies of Acer ne- Steppic: Crimean Mountains 0.00 32 gundo, Amelannchier spicata, Erigeron canadensis, Prunus All biogeoregions 1.13 2701

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Figure 4: Proportions of kenophytes in geographical groups of relevés (total and in habitats G1.1, G1.2a, G1.2b, G1.4. G1.6a, G1.6b, G1.7a). Slika 4: Delež kenofitov v geografskih skupinah popisov (skupno in v habitatnih tipih G1.1, G1.2a, G1.2b, G1.4. G1.6a, G1.6b, G1.7a).

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Figure 5: Proportions of kenophytes in geographical groups of relevés (habitats G1.8, G1.Aa, G1.Ab. G3.1a, G3.1b, G3.2, G3.4a, G3.4b). Slika 5: Delež kenofitov v geografskih skupinah popisov (habitatni tipi G1.8, G1.Aa, G1.Ab. G3.1a, G3.1b, G3.2, G3.4a, G3.4b).

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Figure 6: Proportions of kenophytes in geographical groups of relevés (habitats G3A, G3.Da) Slika 6: Delež kenofitov v geografskih skupinah popisov (habitatni tipi G3A, G3.Da)

Great Britain (the Atlantic biogeoregion, 3.1%). For co- Discussion niferous forests, these values are 0.0%, 0.4% and 24.8% In general, the distribution of proportions of kenophytes respectively. Low proportions of neophytes were found by different forest types of Ukraine is similar to that of the also in Slovenian forests (0.5% in deciduous and 0.1% same forest types of entire Europe presented in the article in coniferous forests) (Küzmič & Šilc 2017). In Ukraine, by Wagner et al. (2017). Habitats with the highest num- the lowest proportion (0.0%) was found in the Crimean ber of relevés and largest occupied areas (G1.6a, G1.A1, Mountains that are the southernmost part of the country. G3.4a) have close values in Ukraine and Europe. Most This area formally belongs to the Steppic biogeoregion, kenophyte-rich alluvial, moist and swampy eutrophic for- but its vegetation has more in common with that of the ests (G1.1, G1.2a, G1.2b, G1.4) in Ukraine have almost northern parts of the Mediterranean biogeoregion. Ac- twice as high proportions of kenophytes than forests of cording to the floristic data (Bagrikova 2013, Yena 2012), the same types from Europe. The main discrepancies are in Crimea, frequent kenophytes of Ukrainian forests are in thermophilous deciduous forests G1.7a, acidophilous absent or rare, less naturalized and more recently estab- beech forests G1.6b and acidophilous oak forests G1.8. lished. The “true” steppic zone (the flatland part of the A lower frequency of kenophytes in Ukrainian forests of Steppic biogeoregion), on the other hand, has the highest G1.6b can be explained by the fact that this habitat is proportion of kenophytes (3.1%). In the relevés from the rare in flatland areas of Ukraine and that all relevés were Steppic biogeoregion including the Crimean Mountains, sampled in the mountains. The proportion of kenophytes the most frequent forest kenophyte of Ukraine and Eu- in habitat G1.7a from Ukraine is higher than in the same rope Impatiens parviflora is not present. habitat in Europe because in Ukraine this habitat is pre- In order to detect changes in the proportion of keno- sented only by relevés from the flatland part of Ukraine phytes over the last few decades, 1498 older relevés sam- with a high proportion of relevés from urbanized areas. pled in 1950–1989 were analyzed. They correspond to 8 The mean value for all relevés is somewhat higher for habitat types. For most of them, a comparison of propor- Ukraine (1.1%) than for Europe (0.9%). One of the tions of kenophytes cannot be made for two reasons: 1) reasons for this difference may be that Ukrainian relevés absence of relevés of the habitat sampled in one of two were sampled more recently. time periods (only “new” or only “old” data), 2) “old” Some data from literature show that the proportion of and “new” relevés represent different areas. Only habitat non-native species in forests of southern Europe is lower G1.Aa “Carpinus and Quercus mesic deciduous wood- than in forests of more northern areas of the Atlantic and land” has large number of “old” and “new” relevés from the Continental biogeographical regions. According to the the same area. This area comprises central and western results of Chytrý et al. 2008 the proportion of neophytes parts of the forest-steppe subregion and adjacent areas in deciduous and mixed deciduous-coniferous forests of of the deciduous forests region of Western Ukraine. Two Catalonia (the Mediterranean biogeoregion) is much low- variants of selection of “new” relevés from this area were er (0.2%) than in the same group of forests in the Czech used: with and without relevés from Kyiv that have a high Republic (the Continental biogeoregion, 1.0%) and proportion of kenophytes. The results of the comparisons

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Table 4: Comparison of kenophyte proportions in “old” and “new” relevés of habitat G1.Aa “Carpinus and Quercus mesic deciduous woodland”. Tabela 4: Primerjava deležev kenofitov v “starih” in “novih” popisih habitatnega tipa G1.Aa “mezofilni listopadni gozdovi vrst rodov Carpinus in Quercus “.

Number of Mean % of Standard Mann-Whitney relevés kenophytes deviation U test Datasets old new old new old new p-level “Old” and “new” data without relevés from Kyiv 520 429 0.261 0.430 0.879 1.174 0.0163 “Old” and “new” data with relevés from Kyiv 520 481 0.261 0.812 0.879 1.831 0.0000 are in Table 4. The data does not satisfy the assumption other words, the increased proportion of kenophytes is of normality so it was tested with nonparametric Mann- statistically reliable in both variants of comparison. In Whitney U test with adjustment for t[[ies. The null hy- fact, the result is a little less certain because of possible pothesis about equal kenophyte proportions in “old” and differences in data sampling by different researchers. “new” relevés was rejected at the 1.63% significance level when “new” data did not include relevés from Kyiv and Viktor Onyshchenko , https://orcid.org/0000-0001-9079- at the 0.00% level for dataset with relevés from Kyiv. In 7241

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Appendix

Biogeographical region Geographical Year(s) and subregion group of releves All forest habitats All forest type Habitat Latitude Longitude Radius (km) of plots Number % of Mean kenophytes of plots Number in subregion * steppic: Crimean mountains G1.6a Ai-Petri 44,499 34,026 2 2004-2005 9 0,00 32 * steppic: Crimean mountains G1.Aa Anharskyi Pass 44,791 34,341 4 2005 5 0,00 32 * continental: forest-steppe G1.Aa Bernashivka 48,587 27,489 2 2002 1 0,00 657 * continental: deciduous forest G3.4a Bile Ozero 51,506 25,728 4 2004 15 0,00 1276 * alpine * Bukovyna alpine 48,197 25,694 32 1995-2005 86 0,73 530 alpine G1.6a Bukovyna alpine 48,214 25,664 29 1995-2005 59 0,60 530 alpine G1.Aa Bukovyna alpine 48,181 25,734 25 2000-2005 22 0,88 530 alpine G3.1b Bukovyna alpine 48,214 25,664 29 1995-2005 5 1,68 530 * continental: deciduous forest * Bukovyna deciduous 48,325 25,880 10 2003-2005 21 0,52 1276 continental: deciduous forest G1.6a Bukovyna deciduous 48,317 25,797 5 2003-2005 14 0,52 1276 continental: deciduous forest G1.Aa Bukovyna deciduous 48,325 25,880 10 2004-2005 3 0,00 1276 continental: deciduous forest G1.Ab Bukovyna deciduous 48,317 25,797 10 2003-2005 4 0,93 1276 * continental: forest-steppe * Bukovyna forest-steppe 48,046 25,946 28 2000-2005 21 0,25 657 continental: forest-steppe G1.6a Bukovyna forest-steppe 48,024 25,901 25 2000-2005 7 0,75 657 continental: forest-steppe G1.Aa Bukovyna forest-steppe 48,048 25,969 26 2000-2005 11 0,00 657 continental: forest-steppe G3.1b Bukovyna forest-steppe 48,024 25,901 29 1995-2005 3 0,00 530 * continental: deciduous forest * Cheremskyi 51,526 25,539 5 1998-2001 46 0,00 1276 continental: deciduous forest G3.4a Cheremskyi 51,526 25,539 5 1998-2001 29 0,00 1276 continental: deciduous forest G3.Da Cheremskyi 51,526 25,539 5 1998-2001 9 0,00 1276 continental: deciduous forest G3.A Cheremskyi 51,526 25,539 5 1998-2001 8 0,00 1276 * alpine G3.1a Chornohora 48,076 24,646 13 2002 7 0,00 530 * continental: forest-steppe G1.Aa Chornukhynshchyna 50,206 33,018 9 1997 25 0,00 657 * continental: forest-steppe G1.Aa Chornyi Lis 48,774 32,581 6 2001 41 0,00 657 * alpine * Chyvchyny 47,849 24,838 16 2002, 2003 20 0,21 530 alpine G3.1a Chyvchyny 47,849 24,838 16 2002, 2003 18 0,21 530 alpine G1.6a Chyvchyny 47,849 24,838 16 2002, 2003 2 0,21 530 * steppic: flatland G1.1 Danube delta 45,404 29,634 23 2003 7 10,16 180 * alpine G1.A4 Dubyno 48,544 22,803 5 2016 4 0,00 530 * alpine * Gorgany 48,441 24,344 8 2005 53 0,11 530 alpine G1.2a Gorgany 48,441 24,344 8 2005 5 0,00 530 alpine G1.6a Gorgany 48,441 24,344 8 2005 7 0,84 530 alpine G3.1a Gorgany 48,441 24,344 8 2005 27 0,00 530 alpine G3.1b Gorgany 48,441 24,344 8 2005 9 0,00 530 alpine G3.2 Gorgany 48,441 24,344 8 2005 5 0,00 530 * continental: forest-steppe G1.Aa Hadiach 50,333 34,027 22 2011-2016 40 1,03 657 * continental: deciduous forest * Halych 49,140 24,620 25 2010-2011 23 1,89 1276 continental: deciduous forest G1.1 Halych 49,083 24,667 9 2010-2011 3 7,89 1276 continental: deciduous forest G1.2b Halych 49,182 24,597 5 2010-2011 4 0,00 1276 continental: deciduous forest G1.6a Halych 49,076 24,600 9 2010-2011 6 0,56 1276 continental: deciduous forest G1.Aa Halych 49,163 24,620 11 2010-2011 10 1,64 1276

301 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine

* alpine * Hutsulshchyna 48,335 24,953 21 2006-2008 73 0,75 530 alpine G1.1 Hutsulshchyna 48,339 25,280 5 2008 3 5,34 530 alpine G1.2a Hutsulshchyna 48,335 24,953 21 2006-2008 11 2,10 530 alpine G1.6a Hutsulshchyna 48,335 24,953 21 2006, 2007 20 0,17 530 alpine G1.6b Hutsulshchyna 48,335 24,953 21 2006, 2007 9 0,00 530 alpine G1.Aa Hutsulshchyna 48,335 24,953 21 2006, 2008 18 0,69 530 alpine G3.1a Hutsulshchyna 48,335 24,953 21 2006, 2007 12 0,00 530 * continental: forest-steppe G1.Aa Ichnia 50,719 32,317 28 2004 6 0,00 657 * continental: forest-steppe G1.4 Irdyn 49,496 31,819 3 1996 3 0,00 657 * continental: deciduous forest G1.Aa Ivano-Frankivsk 48,972 24,683 10 2006-2007 7 0,29 1276 * continental: forest-steppe * Kaniv 49,775 31,535 14 1990-1995 53 4,91 657 continental: forest-steppe G1.1 Kaniv 49,719 31,569 2,5 1990-1995 15 16,17 657 continental: forest-steppe G1.7a Kaniv 49,855 31,549 1,7 1990-1995 9 0,00 657 continental: forest-steppe G1.Aa Kaniv 49,717 31,500 4 1990-1995 26 0,67 657 continental: forest-steppe G3.4a Kaniv 49,855 31,549 1,7 1990-1995 3 0,00 657 * steppic: flatland * Kherson 46,483 32,291 54 1999-2010 60 3,13 180 steppic: flatland G1.1 Kherson 46,482 32,676 25 2000-2010 12 4,29 180 steppic: flatland G1.2a Kherson 46,484 32,144 42 1999-2010 16 2,49 180 steppic: flatland G1.2b Kherson 46,484 32,144 42 1999-2010 8 1,39 180 steppic: flatland G1.7a Kherson 46,484 31,911 21 1998-2009 24 3,56 180 * continental: forest-steppe * Khmelnytskyi 49,418 27,052 12 2004 28 1,23 657 continental: forest-steppe G1.7a Khmelnytskyi 49,325 27,007 0,5 2004 3 1,15 657 continental: forest-steppe G1.Aa Khmelnytskyi 49,418 27,052 12 2004 25 1,24 657 * continental: forest-steppe * Khorol 50.0580 33.6073 67 2001-2004 60 2,77 657 continental: forest-steppe G1.1 Khorol 50.0580 33.6073 67 2001-2004 8 8,21 657 continental: forest-steppe G1.2a Khorol 50.0580 33.6073 67 2001-2004 11 5,37 657 continental: forest-steppe G1.4 Khorol 50.0580 33.6073 67 2001-2004 8 1,35 657 continental: forest-steppe G1.Aa Khorol 50.0580 33.6073 67 2003-2004 33 0,93 657 * continental: deciduous forest * Korostyshiv 50,270 29,095 25 2001-2004 23 0,48 1276 continental: deciduous forest G1.7a Korostyshiv 50,270 29,095 25 2001-2004 10 0,42 1276 continental: deciduous forest G1.Aa Korostyshiv 50,270 29,095 25 2001-2004 6 0,00 1276 continental: deciduous forest G1.Ab Korostyshiv 50,270 29,095 25 2004 7 3,65 1276 * continental: forest-steppe G1.7a Kotovsk 47,807 29,493 12 2007 15 2,25 657 * continental: forest-steppe * Kremenchuk 49,053 33,428 9 2000-2005 24 7,85 657 continental: forest-steppe G1.1 Kremenchuk 49,053 33,428 9 2000-2005 17 9,84 657 continental: forest-steppe G1.4 Kremenchuk 49,053 33,428 9 2000-2005 7 3,02 657 * continental: deciduous forest G1.Aa Kremenets 50,074 25,655 2 1999 7 0,00 1276 * continental: deciduous forest * Kyiv west 50,506 30,302 10 2015-2017 48 4,91 1276 continental: deciduous forest G1.7a Kyiv west 50,506 30,302 10 2015-2018 14 4,42 1276 continental: deciduous forest G1.2a Kyiv west 50,506 30,302 10 2015-2019 9 5,83 1276 continental: deciduous forest G1.2b Kyiv west 50,506 30,302 10 2015-2020 9 5,75 1276 continental: deciduous forest G1.4 Kyiv west 50,506 30,302 10 2015-2021 5 2,53 1276 continental: deciduous forest G1.Aa Kyiv west 50,506 30,302 10 2015-2022 3 5,84 1276 continental: deciduous forest G3.4a Kyiv west 50,506 30,302 10 2015-2022 8 6,84 1276 * continental: forest-steppe * Kyiv south 50,302 30,544 23 2003-2017 154 4,35 657 continental: forest-steppe G3.4a Kyiv south 50,291 30,577 9 2003-2017 33 4,71 657 continental: forest-steppe G1.7a Kyiv south 50,333 30,541 8 2008-2014 4 1,99 657 continental: forest-steppe G1.4 Kyiv south 50,333 30,541 8 2003-2015 15 4,42 657

302 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine

continental: forest-steppe G1.Aa Kyiv south 50,333 30,541 8 2003, 2009 53 3,89 657 continental: forest-steppe G1.2a Kyiv south 50,333 30,541 8 2003-2016 7 4,20 657 continental: forest-steppe G1.2b Kyiv south 50,333 30,541 8 2003-2016 42 4,88 657 * continental: forest-steppe G3.4a Kyiv plato 49,941 30,673 60 1996-1991 8 0,00 657 * steppic: Crimean mountains G1.6a Laspi 44,419 33,756 3 2004-2005 5 0,00 32 * continental: deciduous forest * Lviv 49,827 24,025 12 1992-1997 17 2,37 1276 continental: deciduous forest G1.6a Lviv 49,827 24,025 12 1992-1997 10 3,09 1276 continental: deciduous forest G1.Aa Lviv 49,827 24,025 12 1992-1997 7 1,34 1276 * alpine * Mala Uholka 48,262 23,632 6 2005-2006 34 0,08 530 alpine G1.6a Mala Uholka 48,275 23,658 4 2005-2006 31 0,00 530 alpine G1.Ab Mala Uholka 48,245 23,618 4 2016 3 0,93 530 * continental: deciduous forest * Medobory 49,282 26,164 15 1995-1998 80 0,24 1276 continental: deciduous forest G1.6a Medobory 49,282 26,164 15 1995-1998 4 0,00 1276 continental: deciduous forest G1.Aa Medobory 49,282 26,164 15 1995-1998 49 0,36 1276 continental: deciduous forest G1.Ab Medobory 49,282 26,164 15 1995-1998 27 0,07 1276 * alpine * Mukachevo alpine 48,473 22,740 3 2003-2004 20 0,63 530 alpine G1.6a Mukachevo alpine 48,478 22,746 3 2003-2004 10 0,98 530 alpine G1.Aa Mukachevo alpine 48,469 22,740 6 2003-2004 10 0,28 530 * pannonian G1.Aa Mukachevo pannonian 48,399 22,670 3 2003-2004 12 0,23 26 * continental: forest-steppe * Mykhailivka 49,609 31,509 7 1998-2003 17 0,00 657 continental: forest-steppe G1.7a Mykhailivka 49,609 31,509 7 2003 5 0,00 657 continental: forest-steppe G3.4a Mykhailivka 49,609 31,509 7 1998-2003 12 0,00 657 * alpine * Mykulychyn 48,401 24,594 8 2002 14 0,89 530 alpine G1.6a Mykulychyn 48,384 24,574 8 2002-2003 6 2,20 530 alpine G3.1a Mykulychyn 48,403 24,570 6 2002 4 0,00 530 alpine G3.1b Mykulychyn 48,392 24,613 4 2002 4 1,79 530 * continental: deciduous forest G1.Aa Panivetska Dacha 48,619 26,594 3 2004 14 0,30 1276 * continental: deciduous forest * Perebrody 51,689 27,100 12 2004 19 2,16 1276 continental: deciduous forest G1.4 Perebrody 51,689 27,100 12 2004 3 0,00 1276 continental: deciduous forest G3.4a Perebrody 51,689 27,100 12 2004 8 5,13 1276 continental: deciduous forest G3.Da Perebrody 51,689 27,100 12 2004 8 0,00 1276 * continental: forest-steppe G1.7a Poltava 49,552 34,632 1 1997 2 0,00 657 * continental: deciduous forest G1.Aa Radomska Dacha 51,948 32,483 5 2002 28 0,00 1276 * continental: forest-steppe * Ros 49,721 30,403 90 1998-2000 43 3,00 657 continental: forest-steppe G1.1 Ros 49,721 30,403 90 1999-2000 15 6,72 657 continental: forest-steppe G1.4 Ros 49,721 30,403 90 1999-2000 13 1,15 657 continental: forest-steppe G3.4a Ros 49,556 30,885 52 1998-2000 15 0,88 657 * continental: deciduous forest * Roztochia 50,001 23,668 20 1990-2000 179 0,25 1276 continental: deciduous forest G1.2a Roztochia 50,001 23,668 20 1996, 2000 13 0,28 1276 continental: deciduous forest G1.4 Roztochia 50,001 23,668 20 1990-1996 12 0,00 1276 continental: deciduous forest G1.6a Roztochia 50,001 23,668 20 1990-1999 44 0,38 1276 continental: deciduous forest G1.7a Roztochia 50,001 23,668 20 1996, 2000 9 0,00 1276 continental: deciduous forest G1.8 Roztochia 50,001 23,668 20 1990-1998 6 0,00 1276 continental: deciduous forest G1.Aa Roztochia 50,001 23,668 20 1990-2000 40 0,31 1276 continental: deciduous forest G1.Ab Roztochia 50,108 23,701 4 1996, 2004 2 0,00 1276 continental: deciduous forest G3.4a Roztochia 50,001 23,668 20 1990-1998 53 0,21 1276 * continental: deciduous forest G3.A Rys 51,261 25,384 1,5 1997 6 0,00 1276 * steppic: flatland G3.4a Samara 48,656 35,329 17 2003 49 5,17 180

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* continental: forest-steppe * Sestrynivska Dacha 49,756 28,886 3 2003-2004 6 1,01 657 continental: forest-steppe G1.2b Sestrynivska Dacha 49,756 28,886 3 2003-2004 5 0,74 657 continental: forest-steppe G1.Aa Sestrynivska Dacha 49,756 28,886 3 2003-2004 1 2,33 657 * continental: deciduous forest * Shepetivka 50,250 26,908 36 1999-2001 59 0,00 1276 continental: deciduous forest G1.2a Shepetivka 50,250 26,908 36 1999-2001 3 0,00 1276 continental: deciduous forest G1.2b Shepetivka 50,250 26,908 36 1998-2001 6 0,00 1276 continental: deciduous forest G1.4 Shepetivka 50,250 26,908 36 1999-2001 3 0,00 1276 continental: deciduous forest G1.7a Shepetivka 50,250 26,908 36 1999-2001 3 0,00 1276 continental: deciduous forest G1.8 Shepetivka 50,250 26,908 36 1999-2001 4 0,00 1276 continental: deciduous forest G1.Aa Shepetivka 50,250 26,908 36 1999-2001 15 0,00 1276 continental: deciduous forest G3.4a Shepetivka 50,250 26,908 36 1997-2001 6 0,00 1276 continental: deciduous forest G3.Da Shepetivka 50,250 26,908 36 1997-2001 19 0,00 1276 * alpine * Skole 49,055 23,359 17 1990-2001 85 0,14 530 alpine G1.2a Skole 49,055 23,359 17 1990-2001 8 0,66 530 alpine G1.6a Skole 49,055 23,359 17 1990-2001 49 0,14 530 alpine G1.6b Skole 49,055 23,359 17 1990-2001 5 0,00 530 alpine G1.Aa Skole 49,055 23,359 17 1990-2003 9 0,00 530 alpine G1.Ab Skole 49,055 23,359 17 1990-2003 3 0,00 530 alpine G3.1b Skole 49,055 23,359 17 1990-2001 11 0,00 530 * steppic: flatland G1.Aa Snizhne 48,035 38,783 1 2003 11 1,46 180 * continental: deciduous forest G3.4a Somyno 51,414 26,916 8 2004 13 0,00 1276 * continental: deciduous forest * Stara Huta 52,277 33,567 21 2012 176 0,45 1276 continental: deciduous forest G1.1 Stara Huta 52,333 33,343 1 2012 4 7,03 1276 continental: deciduous forest G1.2a Stara Huta 52,297 33,551 13 2012 2 0,00 1276 continental: deciduous forest G1.2b Stara Huta 52,242 33,364 5 2000-2010 16 1,16 1276 continental: deciduous forest G1.4 Stara Huta 52,292 33,541 12 1998-2010 12 0,57 1276 continental: deciduous forest G1.7a Stara Huta 52,277 33,567 21 2001-2012 6 1,87 1276 continental: deciduous forest G1.8 Stara Huta 52,339 33,763 7 1997-2010 16 0,00 1276 continental: deciduous forest G1.Aa Stara Huta 52,328 33,732 5 1997-1999 30 0,00 1276 continental: deciduous forest G3.4a Stara Huta 52,276 33,620 18 1997-2012 72 0,19 1276 continental: deciduous forest G3.Da Stara Huta 52,342 33,698 10 1997-2012 18 0,00 1276 * steppic: Crimean mountains G1.Aa Staryi Krym 45,021 35,072 4 2007 13 0,00 32 * continental: forest-steppe * Sumy 50,678 34,859 55 1996-2002 59 0,12 657 continental: forest-steppe G1.2b Sumy 50,678 34,859 55 1996-2002 5 0,00 657 continental: forest-steppe G1.4 Sumy 50,678 34,859 55 1996-2002 4 0,00 657 continental: forest-steppe G1.7a Sumy 50,678 34,859 55 1996-2002 19 0,38 657 continental: forest-steppe G1.Aa Sumy 50,678 34,859 55 1996-2002 20 0,00 657 continental: forest-steppe G3.4a Sumy 50,678 34,859 55 1996-2002 11 0,00 657 * steppic: flatland * Sviati Hory 48,956 37,644 25 2006 44 0,12 180 steppic: flatland G1.7a Sviati Hory 48,988 37,493 11 2006 3 0,51 180 steppic: flatland G1.Aa Sviati Hory 48,988 37,493 11 2006 24 0,15 180 steppic: flatland G3.4b Sviati Hory 49,029 37,493 7 2001 17 0,00 180 * alpine * Synevyr 48,508 23,705 16 1990-2016 96 0,00 530 alpine G1.2a Synevyr 48,508 23,705 16 1990-2016 5 0,00 530 alpine G1.Ab Synevyr 48,508 23,705 16 1990-2016 2 0,00 530 alpine G1.6a Synevyr 48,508 23,705 16 1990-2016 22 0,00 530 alpine G1.6b Synevyr 48,508 23,705 16 1990-2016 14 0,00 530 alpine G3.1a Synevyr 48,508 23,705 16 1990-2016 49 0,00 530

304 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine

alpine G3.1b Synevyr 48,508 23,705 16 1990-2016 4 0,00 530 * steppic: flatland G1.7a Tarutyno 46,206 29,146 4 2007 4 5,21 180 * continental: deciduous forest G3.A Tkhoryn 51,415 28,317 15 1999 3 0,00 1276 * continental: deciduous forest * Tsuman 50,926 25,884 9 2002-2003 15 0,00 1276 continental: deciduous forest G1.7a Tsuman 50,875 25,965 0,5 2002-2003 3 0,00 1276 continental: deciduous forest G1.Aa Tsuman 50,926 25,884 8 2002-2003 12 0,00 1276 * continental: forest-steppe G1.Aa Uman 48,719 30,243 5 2004-2005 22 1,81 657 * continental: forest-steppe G1.Aa Vasyshchevo 49,826 36,278 1 2006 8 0,00 657 * continental: deciduous forest * Verkhniobuzki Lisy 49,914 24,941 9 2002-2006 387 0,17 1276 continental: deciduous forest G1.6a Verkhniobuzki Lisy 49,914 24,941 9 2002-2005 143 0,11 1276 continental: deciduous forest G1.Aa Verkhniobuzki Lisy 49,914 24,941 9 2002-2006 223 0,17 1276 continental: deciduous forest G1.Ab Verkhniobuzki Lisy 49,914 24,941 9 2002-2005 21 0,48 1276 * pannonian G1.Ab Vynohradiv-Shaian 48,131 23,227 13 2016 14 1,69 26 * alpine * Vyzhnytsia 48,192 25,208 8 1999-2004 38 0,00 530 alpine G1.2a Vyzhnytsia 48,192 25,208 8 2002-2004 3 0,00 530 alpine G1.6a Vyzhnytsia 48,192 25,208 8 1999-2004 11 0,00 530 alpine G1.6b Vyzhnytsia 48,192 25,208 8 2004 9 0,00 530 alpine G1.Ab Vyzhnytsia 48,192 25,208 8 2002 3 0,00 530 alpine G3.1a Vyzhnytsia 48,192 25,208 8 2002 4 0,00 530 alpine G3.1b Vyzhnytsia 48,192 25,208 8 1999-2004 8 0,00 530 * steppic: flatland G1.7a Zaporizki Balky 47,817 35,201 35 2003-2006 5 1,66 180 * continental: deciduous forest G1.7a Zhukiv Khutir 50,483 30,080 2 1996 13 3,68 1276 * continental: forest-steppe G1.Aa Zhuravlivska Dacha 48,597 28,714 4 2003 21 1,70 657 * continental: deciduous forest * Zhytomyrska 50,840 28,334 81 2000-2003 77 0,06 1276 continental: deciduous forest G3.4a Zhytomyrska 50,840 28,334 81 2000-2003 53 0,09 1276 continental: deciduous forest G3.Da Zhytomyrska 50,840 28,334 81 2000-2002 24 0,00 1276

305 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine in 530 530 180 657 657 530 530 530 180 530 657 180 657 657 657 180 1276 1276 1276 1276 1276 1276 1276 Number Number of releves of releves subregion

0,00 29,4 3,64 0,00 5,28 0,00 0,00 0,00 0,00 23,08 39,84 59,07 52,47 71,12 23,66 16,02 51,48 65,68 28,12 11,12 242,55 167,28 100,80 in group Product of Product mean % of kenophytes in group by by in group

number of releves number of releves

European dataset European Mean for for Mean 4,7 1,3 1,8 7,89 0,47 0,95

groups

Median Median for geo - graphical

graphical groups graphical

- geo for Mean

8,41 1,74 1,74 Mean for dataset for Mean 9,13 2,29 3,05 ha) 40,00 100,00 300,00 Area of Area

Ukraine Ukraine

habitat in (thousands

kenophytes Mean % of of % Mean 4,2

0,28 0,00 0,00 2,10 2,49 5,37 5,83 0,00 0,66 0,00 0,00 5,34 7,03 7,89 4,29 8,21 9,84 6,72 0,00 1,39

10,16 16,17

of releves of No. 2 5 7 9 3 8 5 3 7 3 4 3 8 4 8 84 93 95 13 11 16 11 15 12 17 15 2012 2005 2003 2008 2012 Year(s)

2006-2008 1999-2010 2001-2004 2003-2016 2015-2019 1999-2001 1990-2001 1990-2016 2002-2004 2010-2011 1990-1995 2000-2010 2001-2004 2000-2005 1999-2000 2010-2011 1999-2010 1996, 2000 Radius (km) Radius 8 8 8 9 5 9 1 5

21 42 67 10 20 36 17 13 16 23 25 67 90 42 2,5 Longitude

24,953 32,144 33.6073 30,541 30,302 23,668 26,908 23,359 33,551 23,705 25,208 24,344 29,634 24,667 25,280 31,569 32,676 33.6073 33,428 30,403 33,343 24,597 32,144

-tude Lati 48,335 46,484 50.0580 50,333 50,506 50,001 50,250 49,055 52,297 48,508 48,192 48,441 45,404 49,083 48,339 49,719 46,482 50.0580 49,053 49,721 52,333 49,182 46,484

Geographical Geographical of releves group G1.1 G1.2a G1.2b Hutsulshchyna Kherson Khorol south Kyiv west Kyiv Roztochia Shepetivka Skole Huta Stara Synevyr Vyzhnytsia Gorgany Danube delta Danube Halych Halych Hutsulshchyna Kaniv Kherson Khorol Kremenchuk Ros Huta Stara Kherson Habitat type Habitat G1.1 G1.1 G1.1 G1.1 G1.1 G1.1 G1.1 G1.1 G1.1 G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2a G1.2b G1.2b

Biogeographical region Biogeographical alpine steppic: flatland continental: forest-steppe continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest alpine continental: deciduous forest alpine alpine steppic: flatland alpine continental: deciduous forest continental: deciduous forest alpine continental: forest-steppe steppic: flatland continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: deciduous forest steppic: flatland Title string Title * * *

306 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 32 32 657 657 657 530 657 530 530 530 530 657 657 657 657 657 657 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 3,70 0,00 0,00 0,00 7,28 5,25 0,43 5,88 3,33 3,44 0,00 0,00 0,00 2,22 66,3 0,00 0,00 0,00 6,84 0,00 51,75 18,56 35,40 30,90 21,14 12,65 14,95 204,96 0,9 0,3 0,57 0,21 1,19 0,56 1,46 0,34 300,00 650,00 4,88 5,75 0,74 0,00 1,16 0,00 0,21 0,84 0,17 0,00 0,60 0,52 0,75 0,56 0,00 3,09 0,00 0,00 0,00 1,35 3,02 2,53 4,42 1,15 0,00 0,00 0,57 0,00 9 5 6 5 2 7 9 7 6 5 3 3 8 7 5 3 4 85 42 16 20 59 14 10 31 15 13 12 12 459 2005 1996 2004 2003-2016 2015-2020 2003-2004 1998-2001 2000-2010 1996-2002 2004-2005 1995-2005 2003-2005 2000-2005 2010-2011 2004-2005 1992-1997 2005-2006 2001-2004 2000-2005 2015-2021 2003-2015 1999-2000 1990-1996 1999-2001 1998-2010 1996-2002 2002, 2003 2006, 2007 3 5 8 5 8 9 3 4 2 3 9 8 10 36 55 29 25 16 21 12 67 10 12 90 20 36 12 55 30,302 28,886 26,908 33,364 34,859 30,541 25,664 25,797 25,901 24,838 24,344 24,600 24,953 33,756 24,025 23,658 34,026 31,819 33.6073 33,428 30,302 30,541 27,100 30,403 23,668 26,908 33,541 34,859 50,506 49,756 50,250 52,242 50,678 50,333 48,214 48,317 48,024 47,849 48,441 49,076 48,335 44,419 49,827 48,275 44,499 49,496 50.0580 49,053 50,506 50,333 51,689 49,721 50,001 50,250 52,292 50,678 G1.4 G1.6a Sestrynivska Dacha Kyiv west Kyiv Shepetivka Huta Stara Sumy Kyiv south Kyiv Bukovyna alpine Bukovyna deciduous Bukovyna forest-steppe Bukovyna Chyvchyny Gorgany Halych Hutsulshchyna Laspi Lviv Uholka Mala Ai-Petri Irdyn Khorol Kremenchuk west Kyiv south Kyiv Perebrody Ros Roztochia Shepetivka Huta Stara Sumy G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.4 G1.2a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.2b G1.2b G1.2b G1.2b G1.2b continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: forest-steppe continental: forest-steppe alpine continental: deciduous forest continental: forest-steppe alpine alpine continental: deciduous forest alpine mountains steppic: Crimean continental: deciduous forest alpine continental: forest-steppe mountains steppic: Crimean continental: forest-steppe continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: forest-steppe * *

307 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 530 530 530 530 530 657 180 657 657 657 657 657 657 180 180 530 530 530 530 1276 1276 1276 1276 1276 1276 1276 1276 1276 9,8 0,0 0,00 6,81 0,00 0,00 0,00 3,45 4,20 7,96 0,00 0,00 0,00 7,22 1,53 0,00 0,00 0,00 0,00 0,00 13,19 16,90 16,30 85,44 33,75 61,88 11,22 20,84 0,9 0,4 0,00 0,83 0,00 1,51 0,00 1,94 80,00 200,00 0,00 0,98 2,20 0,38 0,14 0,00 0,11 0,00 1,15 2,25 0,00 0,00 0,00 0,51 5,21 0,00 3,56 0,42 4,42 1,99 0,00 1,87 0,38 0,00 0,00 0,00 0,00 0,00 4 6 3 5 2 9 3 4 9 4 3 6 3 9 9 5 37 10 44 49 22 11 15 24 10 14 19 14 151 143 2004 2007 2003 1997 2006 2007 2004 1995-1998 2003-2004 2002-2003 1990-1999 1990-2001 1990-2016 2002-2005 1999-2004 1990-1995 1998-2009 2001-2004 2015-2018 2008-2014 1999-2001 2001-2012 1996-2002 2002-2003 1990-2001 1990-2016 1996, 2000 2006, 2007 3 8 9 8 8 7 1 4 8 20 17 16 15 21 25 12 10 20 36 21 55 11 21 17 16 0,5 0,5 1,7 22,746 24,574 23,668 23,359 23,705 24,941 25,208 26,164 31,911 27,007 29,095 29,493 30,302 30,541 31,509 34,632 23,668 26,908 33,567 34,859 37,493 29,146 25,965 31,549 25,208 24,953 23,359 23,705 48,478 48,384 50,001 49,055 48,508 49,914 48,192 49,282 46,484 49,325 50,270 47,807 50,506 50,333 49,609 49,552 50,001 50,250 52,277 50,678 48,988 46,206 50,875 49,855 48,192 48,335 49,055 48,508 G1.6b G1.7a Mukachevo Mykulychyn Roztochia Skole Synevyr Lisy Verkniobuzki Vyzhnytsia Medobory Kherson Khmelnytskyi Korostyshiv Kotovsk west Kyiv south Kyiv Mykhailivka Poltava Roztochia Shepetivka Huta Stara Sumy Hory Sviati Tarutyno Tsuman Kaniv Vyzhnytsia Hutsulshchyna Skole Synevyr G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.6a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.7a G1.6b G1.6b G1.6b G1.6b alpine alpine continental: deciduous forest alpine alpine continental: deciduous forest alpine continental: deciduous forest steppic: flatland continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: forest-steppe steppic: flatland steppic: flatland continental: deciduous forest alpine continental: forest-steppe alpine alpine alpine * *

308 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 26 32 657 530 657 657 657 657 657 530 657 657 657 657 530 657 180 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 0 0 0 0 31 4,2 2,8 2,76 9,38 0,00 0,00 41,2 0,00 0,00 8,30 0,00 0,00 0,00 0,00 17,40 17,52 30,69 17,42 12,40 16,40 19,36 47,84 206,17 1,4 0,8 0,00 0,28 0,00 0,71 0,00 0,64 800,00 1800,00 0,30 0,23 0,28 0,36 1,34 3,89 5,84 0,00 0,00 0,93 1,24 0,67 0,00 0,69 1,64 1,03 0,00 0,00 0,00 0,00 0,88 0,00 3,68 1,66 0,00 0,00 0,00 0,00 7 3 7 6 6 3 1 5 5 4 6 14 12 10 49 53 33 25 26 18 10 40 41 25 11 22 26 13 16 911 2004 1999 2004 2004 2001 1997 2001 1996 2005 2003-2004 2003-2004 1995-1998 1992-1997 2015-2022 2001-2004 2003-2004 1990-1995 2010-2011 2011-2016 2000-2005 2004-2005 2000-2005 2003-2006 1997-2010 1999-2001 1990-1998 2003, 2009 2006, 2008 3 3 6 8 2 4 6 9 2 2 4 7 15 12 10 25 67 12 28 21 11 22 26 10 22 35 36 20 26,594 22,670 22,740 26,164 24,025 30,541 30,302 25,655 29,095 33.6073 27,052 31,500 32,317 24,953 24,620 34,027 32,581 33,018 25,969 25,880 25,734 27,489 30,080 35,201 34,341 33,763 26,908 23,668 48,619 48,399 48,469 49,282 49,827 50,333 50,506 50,074 50,270 50.0580 49,418 49,717 50,719 48,335 49,163 50,333 48,774 50,206 48,048 48,325 48,181 48,587 50,483 47,817 44,791 52,339 50,250 50,001 Panivetska Dacha Panivetska Mukachevo pannonian Mukachevo Mukachevo alpine Mukachevo Medobory Lviv Kyiv south Kyiv Kyiv west Kyiv Kremenets Korostyshiv Khorol Khmelnytskyi Kaniv Ichnia Hutsulshchyna Halych Hadiach Chornyi Lis Chornukhynshchyna Bukovyna forest-steppe Bukovyna Bukovyna deciduous Bukovyna Bukovyna alpine Bukovyna Bernashivka G1.8 G1.Aa Zhukiv Khutir Zaporizki Balky Zaporizki Anharskyi Pass Stara Huta Stara Shepetivka Roztochia G1.8 G1.8 G1.8 G1.7a G1.7a G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa continental: forest-steppe pannonian alpine continental: deciduous forest continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: forest-steppe alpine continental: deciduous forest continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: deciduous forest alpine continental: deciduous forest steppic: flatland continental: forest-steppe continental: deciduous forest mountains steppic: Crimean continental: deciduous forest continental: deciduous forest * *

309 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 32 26 657 657 657 180 657 530 180 530 530 530 657 530 530 530 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 0 0 3,6 35,7 2,64 0,00 0,00 0,00 0,00 3,72 9,98 0,00 0,00 0,00 2,33 0,00 2,03 2,00 2,78 0,00 3,01 0,00 0,00 38,80 39,82 23,66 16,06 12,40 0,8 0,1 0,07 0,00 0,41 0,03 0,50 0,03 5,00 500,00 1,70 0,17 0,33 1,81 0,00 0,15 0,00 0,00 0,00 1,69 0,00 0,93 1,46 0,48 0,00 0,00 0,00 0,00 2,33 0,00 0,29 0,07 0,31 0,93 0,00 0,43 0,00 0,00 8 3 4 9 2 3 1 2 7 3 7 4 7 21 22 12 24 20 13 14 30 11 21 15 27 40 28 90 223 121 2003 2006 2006 2002 2007 2016 2003 2016 2002 2016 2002 2002-2006 2004-2005 2002-2003 1996-2002 1997-1999 2003-2005 2002-2005 1990-2003 1990-2016 1999-2001 1990-2003 2003-2004 2006-2007 1995-1998 1990-2000 2001-2004 1996, 2004 4 9 1 5 8 8 4 5 5 1 9 3 4 4 5 3 11 55 13 17 16 36 17 10 15 20 25 13 28,714 24,941 36,278 30,243 25,884 37,493 34,859 25,208 35,072 23,227 33,732 25,797 38,783 24,941 23,359 23,705 26,908 23,359 28,886 23,701 24,683 26,164 23,668 23,618 32,483 29,095 22,824 24,646 48,597 49,914 49,826 48,719 50,926 48,988 50,678 48,192 45,021 48,131 52,328 48,317 48,035 49,914 49,055 48,508 50,250 49,055 49,756 50,108 48,972 49,282 50,001 48,245 51,948 50,270 48,565 48,076 Zhuravlivska Dacha Verkniobuzki Lisy Verkniobuzki Vasyshchevo Uman Tsuman Sviati Hory Sviati Sumy Vyzhnytsia Staryi KrymStaryi Vynohradiv-Shaian Stara Huta Stara Bukovyna deciduous Bukovyna Snizhne Verkniobuzki Lisy Verkniobuzki Skole Synevyr Shepetivka Skole Sestrynivska Dacha Roztochia Ivano-Frankivsk Medobory Roztochia Mala Uholka Mala Radomska Dacha Korostyshiv G1.Ab G3.1a Dubyno Chornohora G3.1a G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Aa G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab G1.Ab continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: forest-steppe continental: deciduous forest steppic: flatland continental: forest-steppe alpine steppic: Crimean mountains steppic: Crimean pannonian continental: deciduous forest continental: deciduous forest steppic: flatland continental: deciduous forest alpine alpine continental: deciduous forest alpine continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: deciduous forest alpine continental: deciduous forest continental: deciduous forest alpine alpine * *

310 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 530 530 530 530 530 530 657 657 657 657 657 180 530 657 530 530 530 530 530 530 1276 1276 1276 1276 1276 1276 1276 3,85 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 8,40 0,00 0,00 7,14 0,00 0,00 0,00 0,00 54,72 41,04 13,20 11,13 155,43 253,33 0 0,1 1,4 0,00 0,00 0,05 0,50 0,00 1,45 0,35 0,00 1,41 0,18 130,00 2700,00 0,00 0,00 0,00 0,00 0,21 0,00 5,13 5,17 0,00 0,00 0,00 0,00 6,84 4,71 0,00 0,00 0,88 0,21 0,00 0,00 1,79 0,00 1,68 0,00 0,00 0,00 0,00 5 4 4 8 3 8 8 6 9 4 5 5 3 4 8 44 27 12 18 49 49 13 15 29 33 12 15 53 11 388 2005 2002 2002 2004 2003 2004 2004 2005 2002 2005 1990-2016 1998-2001 1990-1995 2015-2022 2003-2017 1996-1991 1998-2003 1998-2000 1990-1998 1997-2001 1995-2005 2000-2005 1990-2001 1990-2016 1999-2004 2006, 2007 2002, 2003 8 6 8 5 9 7 8 4 8 6 8 8 21 16 16 10 60 12 52 20 17 36 29 25 17 16 1,7 24,344 24,953 24,570 23,705 25,208 24,838 25,539 31,549 30,302 30,577 30,673 31,509 27,100 30,885 23,668 35,329 26,908 26,916 25,728 25,664 25,901 24,344 24,613 23,359 23,705 25,208 24,344 48,441 48,335 48,403 48,508 48,192 47,849 51,526 49,855 50,506 50,291 49,941 49,609 51,689 49,556 50,001 48,656 50,250 51,414 51,506 48,214 48,024 48,441 48,392 49,055 48,508 48,192 48,441 G3.1b G3.2 G3.4a Gorgany Hutsulshchyna Mykulychyn Synevyr Vyzhnytsia Chyvchyny Cheremskyi Kaniv west Kyiv south Kyiv plato Kyiv Mykhailivka Perebrody Ros Roztochia Samara Shepetivka Somyno Bile Ozero Bile Bukovyna alpine Bukovyna Gorgany Bukovyna forest-steppe Bukovyna Gorgany Mykulychyn Skole Synevyr Vyzhnytsia G3.2 G3.1a G3.1a G3.1a G3.1a G3.1a G3.1a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.4a G3.1b G3.1b G3.1b G3.1b G3.1b G3.1b G3.1b alpine alpine alpine alpine alpine alpine continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: forest-steppe continental: forest-steppe continental: deciduous forest continental: forest-steppe continental: deciduous forest steppic: flatland continental: deciduous forest continental: deciduous forest alpine alpine continental: deciduous forest continental: forest-steppe alpine alpine alpine alpine alpine * * *

311 Viktor Onyshchenko 18/2 • 2019, 289–312 Kenophytes in different forest types of Ukraine 657 180 1276 1276 1276 1276 1276 1276 1276 1276 1276 1276 0,00 0,00 0,00 4,98 0,00 0,00 0,00 0,00 0,00 0,00 0,00 13,90 0 0 0,1 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,00 0,50 12,00 500,00 0,00 0,00 0,19 0,00 0,09 0,00 0,00 0,00 0,00 0,00 0,00 0,00 3 6 8 8 9 17 17 78 72 11 53 19 18 24 17 1999 1997 2004 2001 1997-2012 1996-2002 2000-2003 1998-2001 1998-2001 1997-2001 1997-2012 2000-2002 5 5 7 15 55 81 18 12 36 10 81 1,5 28,317 25,384 34,859 28,334 33,620 25,539 27,100 26,908 33,698 28,334 25,539 37,493 51,415 51,261 50,678 50,840 52,276 51,526 51,689 50,250 52,342 50,840 51,526 49,029 Tkhoryn Rys G3.4b G3.A G3.Da Sumy Zhytomyrska Stara Huta Stara Cheremskyi Perebrody Shepetivka Huta Stara Zhytomyrska Cheremskyi Sviati Hory Sviati G3.A G3.A G3.A G3.4a G3.4a G3.4a G3.4b G3.Da G3.Da G3.Da G3.Da G3.Da continental: deciduous forest continental: forest-steppe continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: deciduous forest continental: deciduous forest steppic: flatland continental: deciduous forest continental: deciduous forest * * *

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